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Received 29 November 2000; received in revised form 21 September 2001; accepted 5 November 2001
Abstract
Montane evergreen forests in northern Thailand are high in biodiversity and becoming increasingly fragmented. We studied
fragmentation and wildlife response in two contiguous wildlife sanctuaries. Om Koi still maintained relatively large patches ( >400
ha) with some connectivity while Mae Tuen was comprised mainly of small and isolated patches ( <100 ha). Mae Tuen lost 2640 ha
of montane evergreen forest between 1954 and 1996 compared to a loss of 888 ha in Om Koi. We compared the wildlife between
four forest patches in Mae Tuen, with four in Om Koi finding nine mammals and 89 birds in Mae Tuen and 19 mammals and 119
birds in Om Koi. Om Koi still supports populations of large mammals and frugivorous birds extirpated in Mae Tuen. The results
document the high rate of fragmentation in protected areas that often interact synergistically with other pressures to reduce biodi-
versity. Crown Copyright # 2002 Published by Elsevier Science Ltd. All rights reserved.
Keywords: Thailand; Birds; Mammals; Fragmentation; Hunting; Protected areas
Fig. 2. Location of Om Koi, Mae Tuen and other protected areas in northern Thailand.
was surveyed seven times during site visits spanning human activities, such as hunting (carcasses, poaching
from September 1997 to June 1998. platforms, poachers, gun shots etc), burning and cattle
Bird surveys focused on diurnal species and were grazing and the sites of these disturbances recorded by
undertaken between 07:00–11:00 and 14:00–17:00, when GPS. This evidence helped construct a more qualitative
birds are most active, on each transect by the same understanding and is used to supplement our quantita-
observer when no appreciable rainfall was occurring. tive data in the discussion.
Using the same observer standardized skill and effort
variability in bird identification (Bibby et al., 1992).
Bird species, number of individuals, and perpendicular 4. Results
distance from the transect lines were recorded. The
software ‘‘DISTANCE version 2.0’’ (Laake et al., 1993) 4.1. Landscape structure and configuration
was used for analysis of bird density (birds/ha). Species
abundance was indicated as ‘‘birds/site visit’’ as the Table 2 shows the comparison between the two mon-
amount of data for individual species was not enough tane evergreen landscapes selected for more detailed
for DISTANCE to produce reliable results. t-Tests were study in the two sanctuaries. The two landscapes are
used to test the null hypotheses that there were no sig- approximately the same size (20,481ha OK, 18,530 MT)
nificant differences between species diversity and density and do not differ appreciably in the total amount of
in: (1) forest patches with high and low rates of frag- evergreen forest remaining in 1996 with 3403 ha, in OK,
mentation/human disturbances, and (2) edges and and 2475 in MT. However, they do differ in two
interior zones. The rate of fragmentation/human dis- important respects. The first is the change over time.
turbances was defined as the change in forest areas, Between 1954 and 1996, OK lost 5% of its montane
sizes, shapes, and other forms of developments (e.g. evergreen and MT 50%. Secondly, not only did MT
roads) over time across the studied landscapes. lose about 10 times as much forest, the remaining forest
We recorded all large and medium sized mammals is also much more highly fragmented than OK. One
encountered during the bird transect surveys. However, critical indicator of the difference in fragmentation
the main data source for mammals was track counts. between the sanctuaries is the largest patch index (LPI)
Track counts have been used as indices of abundance representing the percentage of the landscape occupied
for species that are difficult to observe on the assump- by the largest patch. In OK this comprises almost 17%
tion that track counts are related to population size in a of the landscape, but in MT only 4%. In 1954 the cor-
linear, or at least a monotonic, fashion (Wemmer et al., responding figure for MT was 23%. There are a larger
1996). In this study, ‘‘track recording stations’’, areas number of smaller patches of more uniform size in MT
51 m on the ground where litter was cleared and soil than in OK. The total core area is also significantly
tilled to create a soft zone for recording foot prints, lower in MT, with only 35% of its extent in 1954,
were established at 20–25 m intervals for every 100 m whereas OK retained some 86% of total core area
along the transect. There were 40 stations in each patch. since 1954.
Each was re-tilled after the tracks had been recorded. T-
tests were used to test the null hypothesis that there was 4.2. Birds
no significant difference of track abundance in each
mammal species between high and low fragmented/dis- Overall we made 2430 sightings of 149 species, with
turbed landscapes. 1238 sightings of 89 species in Mae Tuen patches and
In addition to the wildlife information recorded above 1192 sightings of 119 species in Om Koi (Table 3). The
comprehensive field notes were also kept on any signs of mean number of species in OK per patch was 63.25
Table 2
Comparisons of montane evergreen forest change between 1954 and 1996 in Om Koi and Mae Tuen Wildlife Sanctuaries
(SE=3.20) with 54 species in Mae Tuen (SE=2.48). were absent from MT, presumably due to the lack of
This difference was not statistically significant and there interior zone in the smaller patches. Ground omnivores
were no significant differences in bird species density in the Pheasant family such as the Kalij pheasant
between patches in OK and MT or between richness (Lophura leucomelana) and rufous-throated partridge
and patch size. (Arborophila rufogularis) were found in OK patches
Abundance was calculated as birds/site visit and with an abundance of 1 and 3 birds/site visit, respec-
based only on species found on transects to ensure the tively. However, only the bar-backed partridge (Arbor-
same search efforts for both sites. Overall almost half of ophila brunneopectus) with the same size and feeding
the bird species at MT (40 out of 89) were found at an guild as the rufous-throated partridge was found, and
abundance of < 1 bird/site visit while in OK this was then with very low abundance (0.57 bird/site visit) in MT.
about one third of the species (42 from 119). Table 4 In terms of feeding guilds the mean abundance of
shows the five most abundant species at each site. The nectarivorous birds using MT patches was significantly
five most abundant species in OK are all montane ever- higher than in OK (Table 5). The streaked spiderhunter
green forest obligates (Lekagul and Round, 1991). In (8 birds/site visit) and little spiderhunter (4.71 birds/site
MT, however, two generalist species, the black bulbul visit) were the most abundant nectarivores in Mae Tuen
(Hypsipetes madagascariensis) and streaked spider- patches. Nectarivores in OK were more diverse but found
hunter (Arachnothera magna) were found among the in lower abundance, with the chestnut-flanked white-eye
five most abundant species. Large frugivorous birds (3.57) and streaked spiderhunter (1.29) ranking first and
such as the brown hornbill (Ptilolaemus tickelli), and second. None of the other feeding guilds showed any
great hornbill (Buceros bicornis) still existed in very low significant difference between the sites (Table 5).
abundance (1.43 and 0.43 birds/site visit respectively) in
OK but neither was found in MT where they appear to 4.3. Mammals
have been extirpated, as they have throughout almost
all the rest of northern Thailand (Poonswad and Kemp, We found a total of 20 species, nine species in Mae
1993; Vidhidharm et al., 1995). Tuen and 19 in OK (Table 6). OK fragments support large
In OK four species of ground insectivores, the slaty- mammals such as elephant (Elephas maximus), Asiatic
bellied tesia (Tesia olivea), pygmy wren-babbler (Pnoe- black bear (Selenartos thibetanus), leopard (Panthera
pyga pusilla), streaked wren-babbler (Napothera brevi- pardus), tiger (Panthera tigris), sambar (Cervus unicolor),
caudata), and dark-sided thrush (Zoothera marginata) and primates such as assamese macaque (Macaca
were found in interior zones, but only one, the dark- assamensis), Phayre’s langur (Presbytis phayrei), and
sided thrush was found in the edge zone. These species white-handed gibbon (Hylobates lar). Except for the
Table 3
Bird and mammal diversity, and bird density in different forest patches in Om Koi (OK) and Mae Tuen (MT)
MT OK
P1 P2 P3 P4 P5 P6 P7 P8
Bird count 289 304 340 305 310 356 241 285
Bird species 50 54 51 61 68 64 54 67
Bird density (birds/ha) 9.51 8.67 8.92 8.10 5.92 7.89 9.06 4.75
Mammal species 7 2 6 5 9 18 7 10
Table 4
Five most abundant species (birds/site visit) in Om Koi and Mae Tuen wildlife sanctuaries
Gray-cheeked fulvetta (Alcippe morrisonia) 11.00 Gray-cheeded fulvetta (Alcippe morrisonia) 16.29
Mountain bulbul (Hypsipetes mcclellandii) 11.00 Black bulbul (Hypsipetes madagascariensis) 12.29
Golden-throated barbet (Megalaima franklinii) 10.86 Mountain bulbul (H. mcclellandii) 10.86
White-tailed leaf-warbler (Phylloscopus davisoni) 8.71 Streaked spiderhunter (Arachnothera magna) 8.00
Black-headed sibia (Heterophasia melanoleuca) 7.57 White-throated bulbul (Criniger flaveolus) 8.00
160 A. Pattanavibool, P. Dearden / Biological Conservation 107 (2002) 155–164
white-handed gibbon none of these animals were found tracks per site visit for all species combined was sig-
in MT. We found smaller size mammals such as the nificantly higher (t=2.74, 12 df, P=0.018) in OK
black giant squirrel (Ratufa bicolor), barking deer (x=271.57 tracks/site visit, SE=46.48) than in MT
(Muntiacus muntjak), wild pig (Sus scrofa) and hog (x=123.43 tracks/site visit, SE=27.78). Barking deer
badger (Arctonyx collaris) at both sites. Although fre- and wild pig tracks came first and second respectively in
quencies of observations were not high enough to abundance at both sites. In OK, elephants were the
undertake statistical tests, the relationship between third most abundant track. Barking deer were sig-
patch size and mammal diversity can be seen in Table 3. nificantly more abundant in OK than MT (t=2.21, 12
Smaller patches show little relationship between size df, P=0.047). No tigers or leopards were recorded at
and diversity, but as patch sizes get larger they provide the track stations.
suitable habitat for an increasing number of species.
Almost twice as many species (18) were found in the
largest patch (P6, 796 ha) when compared with the next 5. Discussion
largest patch (10 species in P8, 462 ha).
5.1. Landscape and patch configuration
4.3.1. Mammal track abundance
The total number of tracks for all species was 886 in These results clearly show that even within the pro-
MT and 2016 in OK (Table 7). The mean number of tected area system the amount of montane evergreen
forest has been severely degraded as a result of agri-
Table 5 cultural activities. Within the selected landscapes in the
Comparisons on bird abundance by feeding guild in montane two sanctuaries, OK lost 5% and MT an astonishing
evergreen forest patches between Om Koi and Mae Tuen wildlife 50% between 1954 and 1996. MT lost more than 10
sanctuaries
times as much montane evergreen as did OK over
Feeding guild Om Koi Mae Tuen t df P Powera this time period. Fragmentation statistics show MT to
x SE x SE have a larger number of smaller patches remaining with
a total core area of only 5% of the landscape, almost
Nectarivore 1.68 0.41 3.64 0.31 3.82 6 0.009* –
Frugivore 10.71 1.68 16.00 1.99 2.03 6 0.088 0.53 half of the 8% found in OK.
Insectivore 29.25 2.34 21.79 2.11 2.34 6 0.058 0.65 Five related causes probably underlie the more rapid
Omnivore 2.10 0.64 2.11 0.50 0.38 6 0.719 0.07 fragmentation in MT. First, there is a paved road run-
a
Statistical power at /=0.05. ning through the south part of the sanctuary that pro-
* Significant difference. vides ready access both for older established villages
Table 6
List of mammal species found in Om Koi and Mae Tuen wildlife sanctuariesa
Total found 9 19
a
F, Found; N/F, not found.
A. Pattanavibool, P. Dearden / Biological Conservation 107 (2002) 155–164 161
Table 7
Relative abundance (average track counts per site visit) of mammal tracks in Mae Tuen and Om Koi, northern Thailand
to get goods out to market and also for new settlers to pared with OK over the last 40 years, but it is difficult to
enter the sanctuary. There are no paved roads through assess other factors such as hunting impacts in any rig-
OK. Second, the accessibility of MT has made the area orous manner. Hence the following discussion is based
prime growing land for temperate crops, particularly on both our quantitative results supplemented by
cabbages, which are sent to Bangkok. Large profits can observations on human impacts and interviews with
be generated by such cash crops (e.g. see Tungittipla- local villagers and sanctuary guards.
korn, 1998). In the 1997/1998 field season we saw many
new pick-up trucks as the market price for cabbage was 5.2. Bird response
high. Fox et al. (1995) also note the huge impact that a
new paved road had on opening up villages to markets Bird species richness was higher in OK than MT.
in northern Thailand. Third, MT has become increas- Several factors could be responsible for this. Patch size
ingly settled by the Hmong ethnic group, who are well is greater in OK and several studies elsewhere (e.g.
known for their skill in cash crop cultivation. Large Bierregaard et al., 1992; Kattan and Alvarez-Lopez,
numbers of illegal and poorly paid immigrants from 1996, Cornelius et al., 2000) have shown relationships
Burma can be witnessed during harvest time carrying between patch size and bird species richness. However,
the cabbages to the waiting trucks of the Hmong. In no statistically significant relationships between patch
contrast, OK is occupied dominantly by the more size and bird species richness, or density were detected.
sedentary and longer established Karen ethnic group, Instead changes in the relative abundance between birds
who have shown less interest in clearing the forests they with different habitat requirements were noted, with an
depend on to grow cash crops for distant markets. The increase in nectarivorous species in the smaller patches
only cabbage growers in OK are the Lahu ethnic group. of MT and a decline in frugivorous species such as the
Most of the ongoing fragmentation is from the expan- brown and great hornbills. Large frugivorous birds
sion of their fields. Fourth, human population numbers require continuous habitat along altitudinal gradients
are also higher in MT than OK, although little is known because fruit availability is variable in time and space,
of growth rates. Fifth, sanctuary authorities have been and tracking these resources involves seasonal move-
more vigilant in OK in their efforts to control defor- ments that cover large areas. Forest fragmentation
estation and other violations. Such vigilance in MT was separates the connection between foraging areas and
arrested when an officer was shot and killed at his guard may severely restrict access to a year-round food supply
post. His surviving colleagues have been less vigilant (Guindon, 1996). Research elsewhere has also suggested
since that time. that frugivorous birds might be more susceptible to
As a result of the interaction among the factors men- fragmentation and hence suitable species for indicating
tioned above there are significant differences between minimum area requirements for conservation. Price et
the two sanctuaries in the extent and rate of fragmenta- al. (1999), for example found that pied imperial pigeons
tion. OK has retained a higher proportion of montane were especially vulnerable to forest loss. Interestingly, in
evergreen forest, there are more large patches and the our study, mountain imperial pigeons (Ducula badia)
average size of patches is larger with more core area were the eighth most abundant species in OK but were
retained as a consequence. not recorded in the heavily fragmented MT.
However, difficulties still remain. All sites are unique Forest destruction also reduces the number of large
and human impacts are multi-dimensional. We have trees which hornbills require for nesting. Reduction of
documented the increased fragmentation in MT com- nesting sites can lead to change in nest competition and
162 A. Pattanavibool, P. Dearden / Biological Conservation 107 (2002) 155–164
reproductive success (Poonswad and Kemp, 1993). (Panthera pardus), and herbivores such as elephants
Large frugivorous birds including three species of (Elephas maximus) and sambars (Cervus unicolor) were
hornbills were eliminated from Hong Kong and Singa- not found in MT. Large mammals, especially top pred-
pore by fragmentation and hunting (Corlett and Turner, ators like tigers and leopards, normally have a large
1997). Hornbills in this study have also suffered from home range. A single female tiger, for example, requires
hunting. Casques of the Great hornbill (Buceros bicor- at least 2000 ha and the male probably at least twice as
nis) were commonly seen in hilltribe villages. Vidhid- large (Sunquist, 1981). Home range sizes of Asian ele-
harm et al. (1995) reported that the species had been phants normally range between 3400 to 80,000 ha
extirpated from the central area of northern Thailand. It (Stuwe et al., 1998). The fragmented forests of Mae
remains to be seen whether this small relict population Tuen, with their intensive agricultural development may
can survive. be too small to support such animals. Laidlaw (2000),
Another factor is the nature of the clearings in the working in protected areas in Malaysia, found that the
two sanctuaries. In OK, the fields tend to be abandoned size of the remaining area of natural forest was the most
and in various stages of regrowth whereas in MT they important factor affecting the nature of mammal com-
tend to be active fields dominated by cabbage growing. munities.
The lower contrast between habitat types in OK could Hunting is another factor behind local extirpations.
help maintain the diversity of birds. Bierregaard and All the ethnic hilltribes have a long history of hunting as
Stouffer (1997) describe a number of primary rain forest both an important source of food and also a part of
birds in an Amazonian forest foraging in adjacent sec- their culture and beliefs (e.g. see Garrett, 1926; Bruver,
ondary forest, and using secondary forest to re-colonize 1973; Dearden, 1995). As populations have increased,
small primary forest fragments nearby. Some evergreen villages have become sedentary and firearms more
forest species, such as the yellow-cheeked tit (Parus spi- effective, there has been simply too great a hunting
lonotus), and black-headed sibia (Heterophasia melano- pressure for wild populations to be sustained. During
leuca), used the regrowth along the patch edges in OK. the course of fieldwork numerous poaching platforms
Some primary forest specialists may avoid the matrix were found as were the carcasses of two banteng (Bos
of modified habitats, while others will adapt to the javanicus), 25% of the herd. Development of the paved
changed landscape (Newmark, 1991; Laurance and road in MT has also contributed to species extirpations
Bierregaard, 1997). However, no primary forest birds through both disturbance and the access it provides for
were found in the cabbage fields, and clearing species, poaching. In highly developed areas such as Singapore,
such as the flavescent (Pycnonotus flavescens), and red- extirpations of large carnivores were caused mainly by
whiskered bulbuls (Pycnonotus jocosus), were never hunting (Corlett and Turner, 1997).
found in the patches far from the edge zones at either Tracks of a male tiger were found at high elevation in
site. Lovejoy et al. (1986) also found that very few sec- OK, which is unusual habitat for tigers as mountainous
ond growth bird species invaded tropical forest patches areas with severe seasonal fluctuations are less optimal
in the Amazon. habitats (Rabinowitz, 1993). The presence of the tiger
The abundance of nectarivorous birds in MT patches could be an indicator of the availability of preferred
was striking and may be related to the abundance of prey species such as sambar, which still exist in OK or it
wild bananas surrounding most patches. These appear could also mean that the tiger is avoiding more pre-
to be one of the main sources of nectar for the birds, ferred habitat at lower elevations where human settle-
and are not as abundant in OK. Nectarivores such as ments and encroachment are rampant. Rabinowitz
hummingbirds generally prove to be less vulnerable to (1993) suggests that hunting, human settlements and
fragmentation than insectivores and frugivores (Bierre- forest degradation from agricultural clearing are the
gaard and Stouffer, 1997). main threats to tiger populations in Thailand. All these
Hunting appears to be a greater threat in MT rather threats are more severe in MT patches where no tigers
than OK. We recorded over double the number of gun- were found.
shots in MT as OK, encountered several poaching par- Patch size and mammal diversity seem to be related in
ties in the former and also came across more carcasses OK but no such trend emerged in MT. The lack of
including the feathers of bird species such as barbets. relationship in MT is probably because the intensity
of human activities overwhelms the effects of patch size
5.3. Mammal response and diversity. As mentioned earlier, many authors (e.g.
Warburton, 1997; Kattan and Alvarez-Lopez, 1996;
The conspicuously low numbers of mammal species Bierregaard and Lovejoy, 1989; Chiarello, 2000) have
in MT patches compared to OK are probably a result reported positive relationship between patch size and
of the combined pressures of fragmentation and hunt- animal diversity. Greater diversity in larger patches can
ing. Large carnivores such as tiger (Panthera tigris), be viewed as the result of having more habitat diversity
Asiatic black bear (Selenartos thibetanus), and leopard (Buckley, 1982) or larger area per se (Simberloff, 1976).
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